Springer Series in Cognitive and Neural Systems Volume1 SeriesEditors VassilisCutsuridis JohnG.Taylor Forfurthervolumes: http://www.springer.com/series/8572 Mario Negrello Invariants of Behavior Constancy and Variability in Neural Systems 123 MarioNegrello OkinawaInstituteofScienceandTechnology Okinawa,Japan [email protected] ISBN978-1-4419-8803-4 e-ISBN978-1-4419-8804-1 DOI10.1007/978-1-4419-8804-1 SpringerNewYorkDordrechtHeidelbergLondon LibraryofCongressControlNumber:2011923227 (cid:2)c SpringerScience+BusinessMedia,LLC2011 Allrightsreserved.Thisworkmaynotbetranslatedorcopiedinwholeorinpartwithoutthewritten permission of the publisher (Springer Science+Business Media, LLC, 233 Spring Street, New York, NY10013, USA),except forbrief excerpts inconnection with reviews orscholarly analysis. Usein connectionwithanyformofinformationstorageandretrieval,electronicadaptation,computersoftware, orbysimilarordissimilarmethodologynowknownorhereafterdevelopedisforbidden. Theuseinthispublicationoftradenames,trademarks,servicemarks,andsimilarterms,eveniftheyare notidentifiedassuch,isnottobetakenasanexpressionofopinionastowhetherornottheyaresubject toproprietaryrights. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface As a cognitive scientist, I realize that in the study of the brain, challenges of epistemologicalnaturematchthoseofempiricalnature.Issueshavetoberesolved on both fronts, and with matching impetus. On the empirical forefront, one has todealwithcomplexitiesofmeasurement,systemidentification,andexperimental paradigms.Ontheepistemologicalside,functionhastobeoutlined,analyzed,local- ized,discussed,explained.Thetwosidesmustbealloyediftheexplanatorysummit istobeachieved. Andasamechanicalengineer,Ifinditdifficulttodispelmynaturaltendencyto seemechanismineverything.Themeaningofthemechanismanalogyinthestudy of organismsand brain function has to be put under adequate light to be service- able.Inthebrain,mechanismsaredynamicalprocesses,highlycontextdependent, that subserve an incredible breadth of organismic behavior. They are evanescent, butreliable.Theyareexpressiblethroughwordsandtheories,butareexhaustedby neither. IfoundnocommondenominatorformyconflictingquandariesuntilIwasintro- ducedtocyberneticsthroughtheworksofvonFoerster,Braitenberg,Wiener,Pask, andAshby,andmorerecently,Varela,Maturana,Pasemann,andBeer.Sopervasive astobeinvisible,cyberneticspercolatedthroughmostofthesciencesinthesecond halfofthepreviouscentury,andis,tomymind,theonlytruemethodofachieving understandingamidthevastamountofknowledgethatallsciencescurrentlyamass. It was not only once or twice that a powerfulepiphanyof minecould be retraced 60yearsinto the past, back to those of the magnificentset of revolutionaryscien- tistswhounderstoodwhatitmeanstounderstand.ItistothemthatIpaydueswhere duesaredue,andinallhumblenessofferthisbookasacyberneticcontributionfrom amechanicalengineer,turnedcognitivescientist,convertedcyberneticist. Some words about the formatof the book are required.Skimming the table of contents,onewillnotice thatI tookthe libertyof dividingitinto two parts.Had I heardsomegoodadvice“keepitshort,”itwouldperhapsindeedhavebeenshorter, more to the point, more concise, and would primarily consist of Part II. But then itwouldnotservea particularpurposeIwanteditto play,whichwas tomakeex- plicitthepathtowardsconclusions.Iwantedittobeacompletesnapshot,orbetter, a long-exposure picture, of the considerations leading to subsequent results. For thatreasonprimarilyI have veryintentionallylaid emphasison epistemologyand v vi Preface theoretical aspects of the problem, hence Part I. This discussion seems to me to beessentialtoframebothproblemsthatcanbefractallycomplexandconclusions thatarestrengthenedwhencontextualized.Ibelievethattheprocessofcommitting complex ideas to paper has paid off immensely, and I hope to be harvesting divi- dendsof the extraeffortin times to come.For that, I mustbegto be excusedof a certainindependenceandperchanceunevennessinthetreatmentoftopics,attimes introductory,attimestechnical,attimescontroversial.Forthatreasonalso,theref- erencesare introducedin the individualchapters. If the readershall indulge me, I believe that he or she may come to appreciate my choice that more (not less) is more. Finally, parts of this book have appeared in print before as journal articles (Chaps.5,8),abookchapter(Chap.7),andanewsletterarticle(Chap.9).Theyhave allundergonesubstantialrevisionin anattemptto deliveramoreunifiedpackage. Finally, in the spirit of an open work, the book can also be said to be ongoing. Theideasin it,especiallytheroleofinvariancein thestudyofbehavior,andcon- stancy and variability in neural systems and behavior, are vast, unexplored, and, I feel, immensely rich. If blinded by my excess enthusiasm I failed to offer the precision and depth the ideas deserve, as a way of apologyI urgeyou to keep on listening.Thereismoretocome. Bonn, Germany MarioNegrello Acknowledgements Amind’sproduceiscontingentonothermindsandcircumstances.Thankyou: Jackie Griego and Judith Degen, for jovially enduring my peculiar rants; Steffen Wischmann,Walter de Back. and RobertMaertin, mentalbrethren;Mark Hallam, worthyopponentandinspiredreviser;PeterKo¨nig,atrainingexample;HodLipson, forhospitality;ChristianRempisandKeyanZahedi,forfriendlyscientificsoftware; Orlei Negrello, my first scientific collaborator; Thomas Christaller, for mindful interaction from Japan to Sankt Augustin; Frank Pasemann, for enthusing and entrustingmewithideas,withadmiration. vii Contents PartI InvariantsofBehavior 1 Introduction................................................................... 3 1.1 VariationandConstancyinNeurobiologyandBehavior............. 3 1.2 ArgumentOutline...................................................... 5 1.2.1 PartI:InvariantsandInvariantsofBehavior................ 5 1.2.2 ModelsandDynamicalSystems............................. 6 1.2.3 PartII:TheOrganismandItsWorld......................... 6 1.2.4 ConvergentEvolutionofBehavioralTraits.................. 7 1.2.5 ModularOrganization........................................ 7 1.2.6 ConceptDuals ................................................ 8 References...................................................................... 9 2 InvariancesinTheory........................................................ 11 2.1 InvariantsinPhysicsandMathematics................................ 11 2.1.1 InvariantsAreThatWhichRemainsWhen SomethingElseChanges..................................... 11 2.1.2 TheoreticalInvariants:Speed of Light, Planck’sConstant............................................. 11 2.1.3 TheoreticalandRelationalInvariants:Energy .............. 12 2.1.4 EmpiricalandRelationalInvariants:LawofGases......... 12 2.1.5 Context-DependentInvariances:Sumof InternalAnglesoftheTriangle .............................. 13 2.1.6 InvariantsPrefigureTheories................................. 14 2.2 InvariantsinBiology................................................... 14 2.2.1 BiologyBorrowedInvariantsfromPhysics andMathematics.............................................. 14 2.2.2 InvariantsinBiologyAreContext-Dependent.............. 14 2.2.3 ThreeExamples:Life,Form,andBehavior................. 15 2.2.4 OntogenyNeedstheEnvironment........................... 17 2.2.5 DevelopmentandBiophysics................................ 17 2.2.6 EvolutionOperatesonOrganisms,NotonGenes........... 18 2.2.7 GenesasInvariantsofSpeciesIdentity...................... 19 ix